The Biomineralization in Social Wasps (Vespinae): the Presence of Statoliths
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Scanning Electron Microscopy Volume 1986 Number 4 Article 37 8-23-1986 The Biomineralization in Social Wasps (Vespinae): The Presence of Statoliths J. S. Ishay Tel Aviv University T. Shimony Tel Aviv University L. Arcan Agriculture Research Organization Follow this and additional works at: https://digitalcommons.usu.edu/electron Part of the Life Sciences Commons Recommended Citation Ishay, J. S.; Shimony, T.; and Arcan, L. (1986) "The Biomineralization in Social Wasps (Vespinae): The Presence of Statoliths," Scanning Electron Microscopy: Vol. 1986 : No. 4 , Article 37. Available at: https://digitalcommons.usu.edu/electron/vol1986/iss4/37 This Article is brought to you for free and open access by the Western Dairy Center at DigitalCommons@USU. It has been accepted for inclusion in Scanning Electron Microscopy by an authorized administrator of DigitalCommons@USU. For more information, please contact [email protected]. SCANNING ELECTRON MICROSCOPY/1986/IV (Pages 1619-1634) 0586/5581/86$.00+0S SEM Inc., AMF O'Hare (Chicago), IL 60666-0507 USA THE BI0MINERALIZATI0NIN SOCIALWASPS (VESPINAE): THE PRESENCEOF STATOLITHS J.S. Ishay(*)l, T. (Benshalom) Shimony1 and L. Arcan2 loepartment of Physiology and Pharmacology, Sackler Faculty of Medicine, Tel Aviv University, Ramat Aviv 69978 2 and Volcani Center, Agriculture Research Organization, Beit Dagan 50250, Israel (Received for publication March 03, 1986, and in revised form August 23, 1986) Abstract Introduction In social wasps granular aggregates occur Only a few members of the Insecta have a mainly in the head region and the body integu developed statocyst: A cuticular nodule behind ment. In the former location they are present the eyes called Palmen's organ is found in the both on the outer and inner side of the frons. head of the adult and larva of Ephemeroptera; On the outer side of the frons, in a groove tra in Dorymyrmex(Hymenoptera) there is a stato versing its center, there is a deep pit sur cyst above the coxa on the metathorax which is rounded by a membrane which forms a cyst-like an invagination of the cuticle, lined with tac sac. In the pit of the hornet Vespa orientalis tile hairs and possessing sand grains within (Linne, 1771), there are aggregates comprislng the cavity. These organs probably function as sllicon (Sl), potassium (K), calclum (Ca) and gravity receptors (Marcus, 1956; Chapman, 1969). iron (Fe), which are arranged in morula-llke In the order Hymenoptera (and others) some fashlon. A granular material is secreted at the species are colored; ln the social wasps base of sensory hairs (sensilla), distributed on (Vespinae), there is a predominantly yellow to both sides of the groove, as well as along their green color found in the cuticula spread over shafts. Aggregates of sillcon and calcium are various parts of the body (Kemper and Dohring, also present on the inner side of the frons. 1967; Guiglla, 1972; Spradbery, 1973; Edwards, The integumental aggregates are comprised 1980), which ls of taxonomlc lmportance: it is of yellow pigment granules. In the abdominal also belleved to serve as a warning signal region these granules, whlch are concentrated (Keeton, 1967). Such pigmented cuticular areas in yellow stripes, assume a quasi-cylindrical in Hymenoptera (as well as the colored wings in shape, measuring 0.6 - 0.8 µmin length and 0.3 Lepidoptera) are formed by an infolding of the -0.4 µmin diameter. Their structure is affect cuticula inside and the pigment aggregates are ed by caste, age, exposure to sunlight, and within granules composed of pteridines within drugs lntroduced through feeding. Feeding on the epidermal cells or above them, unlike the colchlcine induces degeneration of the granules, melanin which is embedded inside the cuticula whereas feedlng on xanthines renders them re (Becker, 1937;; Desclmon, 1965; Harmsen, 1966; fractory to the destructive action of colchl Ziegler and Harmsen, 1969). These pigmented cine. Thelr main metalllc element is K. areas in social wasps have photoconductive as We postulate that 1) the granules on both well as thermoconductive properties (Croitoru sldes of the frons are part of a tympanlc and et al., 1978; !shay and Croitoru, 1978; !shay static organ; 2) the yellow granules (YG) dis et al., 1982). Slmilar, although not identlcal, persed all over the external surface of the body pigmented granules have also been observed in possess properties of organic semiconductors; honeybees (J.S. !shay, unpublished observation). 3) land 2 are piezoelectric, and are involved Static sense organs composed of a hollow both in gravlty orlentation and transposition statocyst surrounded by sensory hairs and con from photo to geotaxis. tainlng statoliths have been discerned in the heads of various species of Vesplnae (!shay et KEYWORDS: Statocysts and Statoliths in Insects; al., 1983; !shay and Shlmony, 1986). The pre Yellow Pigment Granules; Photo- and Thermoelect sent paper provides data on the variety of ric Propertles; Biomineralization; Bioaggregates granules encountered In social wasps, and also containing Si, K, Ca, Fe; Gravity Orientation in speculates on their probable functions. Insects. Materials and Methods (*)Address for correspondence: Jacob S. !shay, Dept. of Physiology and The majority of the observations were made Pharmacology, Sackler Faculty of Medicine, Tel on the adult population (workers, queens Aviv University, Ramat Aviv, Tel Aviv 69978, Israel Phone No. (03) 420 863 1619 !shay, J.S., (Benshalom) Shimony, T., Arcan, L. and males) of the Vespinae species: on Vespa Thln sections were stained wlth l.5°/o orientalis, collected from natural nests uranyl acetate and 0.3°/o lead citrate. around the Tel Aviv area in Israel, and on L Micrographs were obtained by a JEOL 1008 crabro (Linne 175B), Paravespula germanica electron microscope. (Fabricius 1793), Dolichovespula media (Retzius Changes in the size and shape of the yellow l7B3) and~ saxonica (Fabr. 1793) collected pigment granules were evaluated after one of in Germany, as well as many other species of the following treatments or conditions: Vespinae and Polistinae (!shay and Shimony, a) exposure of hornet workers to continuous l9B6). The hornets used for experiments were light for 14 days; b) the effect of age, i.e., maintained in artificial breeding boxes (ABBs) the difference between young hornets (0-24 h), (!shay and Sadeh, 1975) in groups of 5-20, and adults (several weeks old), and 'old' queens kept on a regular diet of 30°/o sucrose solu (one year of age); c) addition to the diet of tion and morsels of meat (for details, see J. caffeine, theophylline, allopurinol, or purine I shay, 1964). alone (in a dosage of about 5 µg/hornet/day) Hornets and wasps were killed by freezing or together with colchicine for two weeks; d) and stored until processing. Specimens were deviation from the normal pattern due to an first observed under light microscopy for pre aberration caused either by excessive yellow liminary location of specific areas. pigment, or by its absence. Scanning electron microscopy (SEM) and energy The photoelectric properties of the yellow dispersive x-ray analysis (EDX) abdominal stripes in situ (i.e., when connected Microstructural investigation as well as to the intact body), or of the separated yellow associated chemical information were carried pigment granules, were examined by observing out by JEOL-35 SEM, Cambridge Stereoscan 180S the change in resistance under illumination and Energy Dispersive X-ray Analysis System - and/or in the dark. This was achieved by PROXANIII equipped with a Si (Li) - Sl2.5-4 exposing the hornet after each measurement to high resolution detector. a low temperature (approximately 0°C), for For microstructural observation, the spec 5-15 min. The change in resistance upon imens were attached to aluminium or copper illumination was measured with an integral stubs with silver paint and sputter-coated by white light from incandescent bulbs of 100 W a 200-300 nm gold layer. and 220 Vat a distance of 20 cm from the Parts of the internal surfaces in front specimen. The change in resistance as a were investigated after fixation for a minimum function of illumination with white light was period of 4 h with 2.5°/o glutaraldehyde in measured in two ways: l) during 15-20 min of O.lM phosphate buffer pH 7.2, dehydrated through irradiation with light (until a saturation graded ethanols, then critical point dried (CPD) level was attained), and 2) by further exposure from liquid carbon dioxide and carbon and/or to a light/dark rhythm every 2 min (i.e., gold-sputtered. intermittent illumination). Control experiments Location of a number of elements presented were made to ascertain that the electrical in the frons area was investigated by x-ray connection, i.e. the silver paint, tungsten microanalysis (EDX) on unfixed as well as on wires or the glass were not in any way photo fixed tissues. Semi-quantitative measurements conductive. (For further details, see !shay were performed on YGs using gelatin blocks as and Croitoru, 1978; I shay et al., 1980). an organic matrix. Methods for evaluating the thermoelectric Stereo pictures at low and high magnifi properties, thermoelectric (Seebeck) effect cation were performed in order to check the and electrical capacitance of the yellow versus proper geometry for x-ray analysis. Minute brown stripes of the hornet and wasp cuticle parts of the above specimens (about 1-2 mm2) were also described earlier (Shimony and !shay, were tailored accordingly under a light micro l98la,b; !shay et al., 1982a; Shimony and scope and attached to a carbon stub by carbon !shay, 1984). In this study individual hornets paint. The EDXwas carried out at l5kV-20kV were attached with glue to a microscope slide accelerating voltage with a 50 and 100 sec between two silver paint electrodes. The (lifetime) counting time for each analysis. yellow stripes on the abdominal segments were Occasionally, a very thin layer of gold, connected to the electrodes with thin tungsten insufficient to obscure the EDXpeaks being wires.